Memoirs: Studies on the Structure, Development, and Physiology of the Nephridia of Oligochaeta VI. The Physiology of Excretion and the Significance of the Enteronephric Type of Nephridial System in Indian Earthworms

Abstract

1. In an earthworm, as in most aquatic invertebrates, urea and ammonia form the main bulk of nitrogenous excretion and there is no trace of uric acid. These excretory products are first formed in the body-wall and gut-wall, pass therefrom into the coelomic fluid and blood, and are thence eliminated to the exterior by the nephridia. In Pheretima urea and ammonia pass out from the nephridia to the exterior either directly through the skin or through the two ends of the gut. 2. Ammonia and urea have been estimated for the first time in the blood, coelomic fluid, and urine of the earthworm. It has been shown that blood is not a mere carrier of oxygen, as Rogers believed, but that it also takes part in carrying urea and ammonia from the body-wall and gut-wall to the nephridia. The blood of the earthworm does not coagulate, indicating absence of fibrinogen. 3. The role of the nephridia in excretion and osmotic regulation has been determined. A comparison of the osmotic pressures of blood, coelomic fluid, and urine shows that the coelomic fluid is hypotonic to the blood, and that urine is markedly hypotonic both to the blood and coelomic fluid. The protein and chloride contents of the blood, coelomic fluid, and urine have been determined with a view to elucidate the differences in their osmotic pressures. It has been found that the urine contains the merest trace of protein, but that the amount of proteins in the blood is about eight times that contained in the plasma of the coelomic fluid. On the contrary, the chloride content of the coelomic fluid-plasma is about 60 per cent, higher than that of the blood-plasma. 4. The part of urine which is excreted from the blood is probably a protein-free filtrate, but the nephridia reabsorb all the proteins passing into them with the coelomic fluid-plasma. Similarly, there is a reabsorption of chlorides on a large scale from the initial nephridial filtrate during its passage through the nephridia. 5. A convenient method has been devised for collecting urine of the earthworm, which has made it possible to collect as much as 25 c.c. of urine in two and a half hours. The rate of excretion of the urine has been determined and it has been found that in an earthworm living in water the outflow of urine in twenty-four hours must be more than 45 per cent, of its body-weight. 6. It seems that an earthworm, when submerged in water, can live like a fresh water animal, and its gut acts as an osmoregulatory organ in addition to the nephridia, but in the soil it lives like a terrestrial animal and the osmo-regulatory function is adequately discharged by the nephridia alone which reabsorb chlorides and proteins, and are also active in the conservation of water. In Pheretima and other earthworms with an enteronephric type of nephridial system, the gut takes a prominent part in reabsorbing the water of the nephridial fluid and conserving water to its maximum extent. 7. The phagocytic section (ciliated middle tube) believed by Schneider to be absent in the nephridia of Pheretima has been shown to be distinctly present; it is also present in the nephridia of Lampito , Eutyphoeus, and Tonoscolex. The brownish yellow granules characteristic of this phagocytic section form a heavy deposit in the septal nephridia of Pheretima posthuma, heavier than that described in Lumbricus. The deposit increases with the age of the earthworm and forms a ‘storage excretory product’. 8. Spectroscopic examination has revealed that these brownish yellow granules, so far believed to be of guanine, are really blood-pigment granules, since a pyridine solution of them shows the two characteristic bands of haemochromogen. With regard to the blood-pigment, the nephridia function as ‘storage kidneys’. 9. The mechanism of nephridial excretion of the earthworm can be analysed into processes of filtration, reabsorption, and chemical transformation. 10. It is probable that the dorsal and ventral phagocytic organs of earthworms are additional excretory organs.